The effect of light availability on leaf area index, biomass production

Plant Soil Environ.
Vol. 59, 2013, No. 12: 543–548
The effect of light availability on leaf area index, biomass
production and plant species composition of park grasslands
in Warsaw
P. Dąbrowski1, B. Pawluśkiewicz1, H.M. Kalaji2, A.H. Baczewska3
1Department
of Environmental Improvement, Warsaw University of Life Sciences SGGW,
Warsaw, Poland
2Department of Plant Physiology, Warsaw University of Life Sciences SGGW,
Warsaw, Poland
3Polish Academy of Sciences Botanical Garden – Center for Biological Diversity
Conservation in Powsin, Warsaw, Poland
ABSTRACT
How light conditions affect development of park grasslands is a question that has not been satisfactory addressed.
The aim of this study was therefore determination of the level to which unfavorable light conditions influence
grassy parks area and relationships between parameters which determine state of turf grasses. Researches were
conducted in two parks in Warsaw, in various light conditions and included measurement of: leaf density, sward
height, leaf area index (LAI), and botanical composition of the communities. The leaf density of shaded areas did
not exceed 70%. LAI value varied from 0.5 to 0.9-fold lower than in the areas in half-shade and in sun. The participation of basic lawn species at Skaryszewski Park was higher under shade, while at Łazienki Królewskie was higher
in full-sunlight areas. The state of tested grassy areas in limited solar radiation does not satisfy the requirements of
recreational and representational functions. The development processes of vegetation coverage were inhibited at
the sites of lower solar radiation. LAI was influenced by both leaf coverage and sward height. Agrostis stolonifera
and Poa trivialis may be recommended to create grass areas under limited solar radiation.
Keywords: green infrastructure; park habitat conditions; urban green area; environmental improvement; leaf density of sward
Park grass communities have many functions in
urban areas (Potchter et al. 2006, Tlustoš et al. 2006,
Godefroid et al. 2007). To obtain park areas with
high quality is particularly difficult task, mainly
due to the variation in light conditions. One of the
main responses of plant growing in limited access
to light is to create the elongated tills and less leaf
coverage and amount of shoots per area (Cavagnaro
et al. 2007). Leaf area index (LAI) is one of the most
important of plant canopy parameters (Yang et al.
2012). Unfortunately, at the moment there is no
literature for the use of LAI ratio as an indicator
for assessing the degradation degree of grassland
in urban parks. There are also no studies considering relationships of this parameter with others and
also defining state of park grasslands, such as leaf
density per area or shoot high. So far, park lawns
were rarely studied ecosystems (Bertoncini et al.
2012). The aim of this study was to answer following research questions: (1) How do unfavorable
light conditions influence the development of the
grassy parks area with decorative and recreational
functions?; (2) Is there any correlation between
LAI and leaf coverage and sward height?, and (3)
Which species could be most recommended for
renovation of grassy areas?
MATERIAL AND METHODS
The grassy areas located in Museum Łazienki
Królewskie (52°12'N, 21°1'E) and Skaryszewski
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Plant Soil Environ.
Park (52°25'N, 21°15'E) was the chosen location for
this study. The research was conducted in period
2010–2012 in 3 research sites: shaded by trees,
half shade, and in sun. Single plot area was equal
to 1 m 2 (Table 1). The total incoming radiation to
the surface of grass communities was measured by
use of the solar meter TM 206 device (Tenmars,
Inc. Taipei, Taiwan). Measurements were made
3 times a day, once a month throughout growth
season. Soil moisture was defined by a time domine
reflectometry (TDR) technique; in each plot 3
measurements were made. Chemical soil properties were performed according to standards (PNR-04031). Plant available P and K concentrations
were performed by the Egner-Riehm method. The
results were assessed on the basis of the numbers
contained in Polish standards for mineral soils
(PN-R-04022, PN-R-04023). All plots characterized with a neutral or slightly acidic reaction. Soil
characterized with high abundance of digestible
form for plant of potassium but low abundance
of digestible form of phosphorus. Total nitrogen
content was within the norms prescribed for minerals soils.
The following measurement were performed
periodically (every month during growth season):
leaf coverage of sward area (percentage cover of
leaves, three replicates per treatment were made),
sward height (main from 3 points of the plot before cut), leaf area index (measured in 5 replicates
by use of the SunScan analysis system (Delta-T
Device, Ltd. Cambridge, UK) with light sensitive
‘wand’), the surface weight of cut sward dry matter (over 3 cm) and its botanical composition by
botanical-weight analyses performed three times
during growing seasons: May, July and September.
Collected data were evaluated by the ANOVA
model and by the Fischer’ test as a post hoc at a 0.05
confidence level. All analyses were performed in
Statistica 10.0 program (StatSoft, Inc. Tulsa, USA).
RESULTS AND DISCUSSION
The light conditions significantly affected leaves
coverage of lawns in analyzed parks (Table 2). The
leaves coverage in shady conditions was similar
during the whole study period and was equal to 63%
in Łazienki Królewskie and 71% in Skaryszewski
Park. This parameter was much higher under
partial shading conditions (82–88%). Even better
leaves coverage (95–98%) was found on the lawn in
full sun. Many authors reported that plant growing in a limited access to light formed longer but
Table 1. Characteristics of the habitat research sites
Łazienki Królewskie
GPS localization
Skaryszewski Park
shade
half-shade
sun
shade
half-shade
sun
35.8099''N,
16.3472''E
29.3795''N,
39.2822''E
28,4748''N,
38.8090''E
35.8099''N,
16.8493''E
36.6678''N,
15.6327''E
59687''N,
34721''E
Total irradiance (W/m 2)
Spring
163
230
507
171
373
499
Summer
127
199
564
137
149
530
Autumn
62
99
371
64
127
325
Spring
25
27
28
25
28
32
Summer
24
35
29
23
27
29
Autumn
18
25
26
15
24
26
Soil moisture (%)
Chemical properties of soils
pHKCL
7.2
7.3
7.4
6.4
6.4
6.3
N (%)
0.339
0.441
0.460
0.281
0.297
0.310
4.7
4.3
4.7
3.9
4.6
3.7
25.7
12.0
12.1
18.8
24.7
25.2
P
K
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(mg × 100 g
DM)
Plant Soil Environ.
Vol. 59, 2013, No. 12: 543–548
Table 2. The leaves coverage and sward height at research areas ± SD
Łazienki Królewskie
shade
half-shade
Skaryszewski Park
sun
shade
half-shade
sun
Leaves coverage on research areas (%)
Spring
62 ± 8 a
86 ± 5 b
98 ± 2 c
69 ± 4 a
86 ± 4 b
97 ± 2 c
Summer
63 ± 8 a
87 ± 5 b
98 ± 2 c
68 ± 2 a
87 ± 4 b
97 ± 2 c
Fall
63 ± 2 a
85 ± 3 b
95 ± 2 c
72 ± 3 a
87 ± 4 b
98 ± 2 c
Sward height (mm)
May
73 ± 9 a
107 ± 9 b
113 ± 6 b
83 ± 7 a
117 ± 6 b
136 ± 8 c
June
102 ± 8 a
116 ± 8 a
120 ± 7 b
102 ± 9 a
124 ± 5 b
144 ± 4 c
July
102 ± 5 a
117 ± 8 b
126 ± 7 c
105 ± 7 a
134 ± 5 b
153 ± 6 c
August
95 ± 10 a
120 ± 7 b
119 ± 9 b
105 ± 9
142 ± 8 b
144 ± 6 b
September
94 ± 6 a
104 ± 5 b
108 ± 8 b
102 ± 5 a
124 ± 5 b
137 ± 5 c
October
93 ± 10 a
116 ± 5 b
126 ± 6 c
98 ± 9 ab
104 ± 5 b
93 ± 10 a
The average for the 2010–2012 period. The means for each park in a season marked by the same letter do not
differ significantly
fewer shoots and had less developed root systems
(Repková et al. 2009, Xu et al. 2010).
In May, plant regrowth was considerably slower
in shaded areas than in partial shade and in sun.
The range of sward height in shade was on average
by 30% lower than in half shade and by 37% lower
than in full sun. In the next part of growth season,
differences between height of sward growing in
shade and in sunlight were lower. In Skaryszewski
Park, height of sward amounted to respectively
21% and 28%, and in Łazienki Królewskie – to 13%
and 25%. In Skaryszewski Park significant differences in sward height in half shade and in full sun
Łazienki Krolewskie
3.5
LAI (m2/m2)
3
Sun
Halfshade
Shade
throughout the growing season were also found,
while in Łazienki Królewskie they were observed
only in June and July (Table 2). These results support those obtained by Vargas et al. (2002).
In Łazienki Królewskie, under shade, LAI values
varied from 0.6 to 0.9 (Figure 1); in half-shade they
varied from 1.2 to 1.9; LAI highest values were
found under sun (2 to 2.8). Significant differences
between sun and half-shade were observed from
July and between sun and shade were throughout
the season. Trend of variability was well accentuated within the growing season under sun. In
Skaryszewski Park it was also found that the lowest
3.5
Skaryszewski Park
3
2.5
2.5
2
2
1.5
1.5
1
1
0.5
0.5
0
0
Figure 1. The relationship between the leaf area index (LAI) ratio and the light conditions during the growing
season. The averages for 2010–2011 period
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Plant Soil Environ.
Table 3. The botanical composition and aboveground biomass of sward (% dry matter) ± SD
Łazienki Królewskie Park
shade
Species
Lolium perenne
summer
autumn
spring
summer
autumn
spring
summer
autumn
13.9 ± 4.1a
0.7 ± 0.1a
22.8 ± 6.1a
0
0
0.6 ± 0.1b
0.1 ± 0b
0.3 ± 0b
0
1.6a
0.3a
2.8a
0
0.3b
0b
0
4.2 ±
Poa pratensis
5.3 ± 0.8a
4.0 ± 0.5a
6.1a
20.1 ±
4.3a
Poa annua
4.3 ± 0.3a
0
Agrostis
stolonifera
2.4 ± 1.2a 55.3 ± 10.6a
Grasses total
23.1 ±
1.4 ±
11.1 ±
36 ±
8.1a
0.9 ± 0.1a
75.4
16.1 ± 2.8a
1.9 ± 0.3a
0.7 ± 0.4
Bellis perennis
3.9 ± 0.8a
0.3 ± 0.1a
Plantago major
0
0
20.0
1.2
Trifolium repens
Others less
than 5%
Total
Aboveground
biomass
(g DM/m 2)
0
3.6 ±
26.8 ± 10.1 12.7 ± 3.1
100
11.2 ± 2.4a
100
12 ± 11.2a
15.6 ±
4.2b
46.3 ±
10.8b
1.1 ± 0.2b 6.1 ± 0.7b
53.9
70.2
2.3 ± 0.4b 1.9 ± 0.2a
12.7 ± 3.6a 35.2 ± 8.1b 3.6 ± 0.6a
0.4 ± 0.1 15.1 ± 2.8a
0
1.0a
1.1 ±
1.0 ± 0.3b
12.5 ±
10.2b
3.6 ± 0.7b
0.3 ± 0.1a 27.2 ± 8.4b 12.3 ± 3.2b 53.2 ± 12.9b
81.5
Forbs total
0
4.3 ± 1.0a 10.0 ± 4.2b 5.5 ± 2.3a
53.2
Taraxacum
officin.
sun
spring
Festuca rubra
Poa trivialis
half-shade
13.4
9.0 ±
3.8a
37.9
1.4 ±
20.6
0.5a
2.4 ±
0.8a
2.2 ± 0.4
6.8 ± 0.3
6.8 ± 0.5
100
100
100
72
0
0.1 ±
20.9 ± 5.8c
0.7c
5.1 ±
13.8 ± 4.7c
44.9
0.3 ± 0.1c
1.1 ± 0.2b
1.7 ± 0.4a
9.8 ±
3.1a
13.4 ± 5.6
100
7.7 ± 2.7a 14.5 ± 5.7a 27.3 ± 7.9b 17.5 ± 0.8a
5.6 ± 1.2b
2.4 ± 0.2c 16.0 ± 3.9c
2.0
42.7 ±
77.7
54.5
0b
+
0.9 ± 0.5a 18.3 ± 7.3c
3.8 ± 0.5b
0
4.8
1.5 ± 0.4c
5.0 ± 2.4a 70.1 ± 20.1c 8.0 ± 1.9c
3.7 ± 0.6
0
3.3 ± 0.2a 24.9 ± 6.1c
16.7b
4.7
14.2 ±
0.4 ± 0.1
18.7
5.1b
23.7 ± 8.3b
10.4 ± 3.9
3.4 ± 0.7
3.1 ± 0.8
100
100
100
27.1 ± 8.1b 20.2 ± 3.9b 26.6 ± 8.8b
Averages for the 2011–2012 period. The means (within each light condition) marked by the same letter do not
differ significantly
LAI values were measured in the shade (0.6 to 0.8),
additionally in this positions seasonal changes were
not observed. In half-shade they ranged from 0.8
to 1.9. The significantly highest values of LAI were
measured in full light conditions (1.8 to 2.8). The
changes of LAI during the growing season were
also observed by many authors. Vargas et al. (2002)
concluded that the reduction of LAI was observed
at the end of August. Li et al. (2010) concluded that
LAI initially increased with increasing shading, but
decreased rapidly with time. Research conducted
by Lambert et al. (1999) showed a significant increase in LAI grassland during one spring month.
Authors correlated this phenomenon with nutrient
level and water supply (in spring time, they were
close to optimum), but our research shown that
biomass did not developed properly in unfavorable
light conditions.
In Łazienki Królewskie the correlation coefficient between LAI and leaves coverage was 0.91,
546
and between LAI and sward height it was equal
to 0.84. In Skaryszewski Park the correlation coefficient between LAI and leaves coverage was
equal to 0.94 and, between LAI and sward height
it was equal to 0.95. According to Lambert et al.
(1999) and Ramirez-Garcia et al. (2012) LAI value
for vascular plant species could be affected by:
number of tillers, number of leaves per tiller and
average area of leaves.
The impact of the solar radiation incoming to
grass communities depends on the part of season
(Tables 3 and 4). In Łazienki Królewskie in spring
the masses of sward growing under shade and
half-shade were similar, but in full sun it was twice
higher. The largest biomass of sward in summer
was in half-shade. It was twice as high as in the
shade and full sun. In autumn, most sunny surfaces
characterized themselves by higher biomass, but
mass growing in partial shade was significantly
higher than in the shade. In Skaryszewski Park,
Plant Soil Environ.
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Table 4. The botanical composition and aboveground mass of sward (% dry matter) ± SD
Skaryszewski Park
shade
Species
half-shade
sun
spring
summer
autumn
spring
summer
autumn
Lolium perenne
0
3.7 ± 1.6a
0
9.2 ± 2.6b
7.5 ± 3.8a
16.5 ± 4.3a 16.1 ± 4.9c 15.3 ± 6.8b 12.9 ± 4.7a
Festuca rubra
0
0
0
0
0.1 ± 0a
Poa pratensis
22.1 ± 8.6a 43.2 ± 20.1a 31.4 ± 10.6a 3.2 ± 1.5b
Poa trivialis
26.9 ± 11.8a 13.4 ± 3.1a 35.1 ± 5.7a
Poa annua
11.7 ± 5.6a
Agrostis
stolonifera
14.9 ± 4.2 18.0 ± 6.1
Grasses total
77.6
Ranunculus
repens
2.0 ± 0.4a
Glechoma
hederacea
Plantago major
Forbs total
Trifolium repens
Others less
than 5%
Total
Aboveground
biomass
(g DM/m 2)
1.9 ± 0.1a
80.2
2.0 ± 0.4b
9.3 ± 3.7b 13.4 ± 4.5a
2.2 ± 0.8a 56.7 ± 15.6b 13.3 ± 5.2b
27.6 ± 4.8
96.3
3.3 ± 2.1
81.7
11.5 ± 4.3
45.8
0
4.2 ± 2.8b
spring
summer
autumn
0.5 ± 0.2
0.2 ± 0.1a 12.0 ± 5.5
2.4 ± 1.8b
5.7 ± 1.4c
1.8 ± 0.4b
17.2 ± 6.2b 28.8 ± 9.8a 13.4 ± 2.3a 16.2 ± 4.2b
0.7 ± 0.3a
23.9 ± 7.8
46.0
3.6 ± 0.5c
0.1 ± 0c
+
19.0 ± 13.8 31.7 ± 14.2 28.2 ± 13.3
70.4
31.7
42.9
0.1 ± 0
2.9 ± 1.8a 14.4 ± 8a
6.2 ± 5.9
1.7 ± 0.8a
1.1 ± 0.5b
6.3 ± 2.8
4.5 ± 1.5a 10.9 ± 2.3a
0.8 ± 0.2a
1.9 ± 0.4b
1.5 ± 0.3b
0.3 ± 0.1a
0.6 ± 0.2c
0.2 ± 0.1c
0.5 ± 0.1a
1.3 ± 0.9a
0.6 ± 0.1a
0.2 ± 0.1a
3.1 ± 1.1b 12.2 ± 1.6b
8.6 ± 3.1b
4.4 ± 1.2b
0
1.1 ± 0.6a
11.5
1.1
7.8
0
0
0.2 ±
0.1a
14.6 ± 4.2
8.1 ± 3.2
100
100
0.2 ±
0.1a
2.4 ± 0.5
100
7.2
+
10.4 ± 4.8
100
28.1
12.1 ±
5.8b
15.1
11.8 ±
3.8b
6.7
4.7 ± 3.1
1.3
19.9 ±
7.4b
7.9
6.8 ± 2.4 c
12 ± 2.4
10.6 ± 2.8
18.2 ± 6.8
12.4 ± 5.3
14.2 ± 8.8
100
100
100
100
100
12.9 ± 3.9a 17.6 ± 11.1a 15.7 ± 14.3a 13.1 ± 3.4a 18.6 ± 10.3a 15.6 ± 8.6a 18.61 ± 6.2a 43.9 ± 12.3b18.2 ± 10.4a
Averages for 2011–2012 period. The mean (within each light condition) marked by the same letter do not differ
significantly
significant differences between the mass of the
sward on areas with different treatment were observed only during the summer.
In Łazienki Królewskie the grasses of the great
influence on sward were creeping species, such as:
Agrostis stolonifera L., Poa trivialis L., Poa pratensis
L., and Lolium perenne L. (Table 3). Poa trivialis
grew most efficiently under full shade and halfshade. During spring and autumn, participation of
this species on shaded areas was the greatest of all
grasses – 26% on average. In half-shade Poa trivialis was the dominant species in summer. Agrostis
stolonifera expanded most efficiently under halfshade and full sun. On half-shaded areas this species
had the largest share in sward in spring (27%) and
autumn (53%). Agrostis stolonifera greatest growth
was noted under full sun conditions during summer
time (70%) and shade (55%). Poa pratensis and Poa
annua during spring and autumn grew best under
full sun conditions. In the summer the shares of
these species were low and did not depend on light
conditions. Lolium perenne expanded the best in
the shade during spring and autumn.
In Skaryszewski Park Poa trivialis, Poa pratensis
and Agrostis stolonifera grew best under shade,
especially during spring and autumn (Table 4).
The share of these species was equal to almost
80% of sward. In half-shade in spring Poa annua
dominated, but in the summer and autumn Agrostis
stolonifera and Poa trivialis dominated. In full
sun Agrostis stolonifera dominated in summer.
Poa trivialis and Lolium perenne were also the
species well developing in those conditions. On
the shaded areas smaller density of leaves and the
presence of species undesirable in recreation lawns
were observed (Bertoncini et al. 2012). Research
conducted at Warsaw lawns by Wysocki (1998)
confirmed a significant share of monocotyledon547
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Plant Soil Environ.
ous plants. Festuca rubra, Lolium perenne and Poa
pratensis are characterized by high sward coverage. The share of Poa trivialis was insignificant.
In conclusion, the state of tested grassy areas
under limited solar radiation in both parks does
not satisfy the requirements of recreation and
representational functions, mainly due to insufficient soil cover and small participation of mains
species, such as Lolium perenne, Poa pratensis
and Festuca rubra. There is a lot of factors, which
can influence on grassy areas, such as fertilizer
application (Hejcman et al. 2007). However, the
nutrient content in the soil at all positions was
similar. The development processes of vegetation
coverage were stopped at the sites of lower solar
radiation, which are evidenced by a decreasing in
the LAI values and aboveground biomass. The high
correlation coefficient between LAI and leaves
coverage and between LAI and sward height were
found in both parks. However, future researches
should be extended to find out which of parameters of aboveground biomass affects value of the
LAI. Agrostis stolonifera and Poa travilis may be
recommended to create grass areas under limited
solar radiation because of large participation of
that species in sward under limited solar radiation.
Hejcman M., Klaudisová M., Schellberg J., Honsová D. (2007):
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Received on February 24, 2013
Accepted on October 31, 2013
Corresponding author:
Prof. Hazem M. Kalaji, Warsaw University of Life Sciences SGGW, Faculty of Agriculture and Biology,
Department of Plant Physiology, Nowoursynowska 159, 02-776 Warsaw, Poland
e-mail: [email protected]
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